('Mitrastemoneae'). Maki- No's Initial Use in 1909 of This Name (As 'Mitrastemmaeae') Is Invalid As No Descrip- Tion in Either Japanese Or English Was Given (Dr

BLUMEA 38 (1993) 221-229

A revision of Mitrastema (Rafflesiaceae)

W. Meijer & J.F. Veldkamp

Summary

Makino called Mitrastemon has Mitrastema (Rafflesiaceae), usually incorrectly Makino, two spe-

cies, one in Southeast Asia and one in Central and South America.

Introduction

Mitrastema was described by Makino (1909) on M. (‘Mitrastemma’) yamamotoi

Makino. He intended to name the genus after the mitre-shaped staminal tube, as

becomes evident when in 1911 he changed the name to Mitrastemon.This change is

On the other Stearn not allowed and Mitrastemon is a superfluous name. hand, as

(1973: 519) has pointed out, the Greek words 'sterna' (thread, stamen) and 'stemma'

confused, ‘Mitrastemma’ be (garland or wreath) are easily and so may regarded as an orthographic error to be corrected under Art. 73.1 (Exx. 2, 3). ‘Mitrastemon’ or

‘Mitrastemma’ have been adopted in laterpublications, and even by the International

Code of Botanical Nomenclature, where the name Mitrastemonaceae, invalid because it is based on an invalid name contrary to Art. 18.1, is conserved (1988: 104).

Since its description Mitrastema yamamotoi has become one of the most widely

of the Rafflesiaceae. Watanabe's investigated taxa Especially papers on its morphol-

and of devoted it is still ogy biology (1933-1937) are masterpieces study; yet not clear what the precise taxonomic position of the genus is and whether it really belongs to the Rafflesiaceae.

Some authors have regarded it as a distinct family, Mitrastemonaceae Makino

(1911) (nom. cons.) (e.g. Cronquist, 1981; 'Mitrastemmataceae', Mabberley, 1987),

treatedit Rafflesiaceae tribe others have as Mitrastemateae ('Mitrastemoneae'). Maki- no's initial use in 1909 of this name (as 'Mitrastemmaeae') is invalid as no description in either Japanese or English was given (Dr. J.N. Westerhoven, Hirosaki, oral

validated comm.). It was by Hayata (1913) ('Mitrastemoneae').

Makino it distinct order Mitrastemonales (1911: 252) even proposed to represent a next to the Nepenthales, but he was apparently never followed in this. Although he called it 'series Mitrastemonales' contrary to Art. 33.4 the name seems valid, if one

that 'series' here is the latinizationof the German accepts 'Reihe' of Art. 17.2.

Cronquist, hedging his bets, wrote “‘Mitrastemon’ has often been included in the

Rafflesiaceae, to which it is evidently related. It is more primitive than the Rafflesia-

!) School of Biological Sciences, University of Kentucky, Lexington, KY 40506, U.S.A. 2 ) Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands. 222 BLUMEA Vol. 38. No. 1, 1993

ceae in its hypogynous, perfect flowers, and its dome-like synandrium is more near-

ly comparable to that of Prosopanche, in the Hydnoraceae, than to anything in the

Rafflesiaceae. WhetherMitrastemon should be included in the Rafflesiaceae or kept

is of taste." as a separate family largely a matter

To us Hayata (1913) has given overwhelming evidence that the genus belongs

used the extended Watanabe to Rafflesiaceae. He following arguments, by (1933-

1937) in great detail:

characteristics different from from 1) The endophytic body shows no those known

the Rafflesiaceaeproper.

2) Bisexual flowers occur in Rhizanthes zippelii (Blume) Spach and Bdallophyton

oxylepis (B.L. Rob.) Harms.

3) The tubularandroecium is not completely unique to Mitrastema. In some species

the also form tube of the East African Berlinianche (Harms) Vattimo stamens a

Vattimo around the pistil, e.g. in B. aethiopica (Welw.) and B. holtzii (Engl.)

Vattimo.

4) The superior ovary is an insufficient reason to exclude Mitrastema from the Raf-

families well flesiaceae. In many other genera with superior as as inferior ovaries

have been admitted. Thus, not surprisingly, semi-inferiorovaries are known in

of in blanchetii some species Pilostyles Guill., e.g. P. R.Br. (Gardner, 1844), P. sessilis Rose (1909), P. thurberi A.Gray (Gray, 1854), and P. ulei Solms-

Laub. (Harms, 1935), at least in the female flowers.

5) All the detailedstructures of the embryology, ovules, and seeds of Mitrastema

agree well with those known for other species of Rafflesiaceae proper.

The multibranched, multifloweredinflorescences of Cytinus L. and Bdallophyton

Eichl. and that these the closest seem primitive so suggest may represent genera to

chlorophyllous ancestors of Rafflesiaceae. All other genera are single-flowered,

though especially in Apodanthes Poit. and Pilostyles it might be considered that the

inflorescences buried inside the host and that flowers are their pop up through the bark. In Mitrastema the flowers are still very gregarious, but they have become less

so in Rafflesia Blume, Rhizanthes Dumort., and Sapria Griff.

Mitrastema also shares withother genera of the family the unicellularovary with

parietal placentas.

Watanabe (1936c) concludedthat the stamen tube is derived from 15-20 (average

16) extrorse anthers, each originally with 4 pollen sacs and a connective extension.

It is pushed off after the pollen has been shed by the growth of the pistil, ensuring

cross-fertilization (the anthers are on the outside of the tube, whilethe stigma is inside

it). Sexual reproduction is apparently obligatory (Watanabe, 1934a) but not always

very successful. At higher latitudes the number of fertile fruits rapidly decreases.

He mentioned (1933a) that ripe fruits always become infested by a fungus. Its

concentrated the chlamydospores are around micropylar area. He suggested (1936d)

that the be fungus might fungicidal to other fungi. It may perhaps also take part in germination.

It is not clear how many species there actually are. Opinions vary between one to five, or even more. W. Meijer & J.F. Veldkamp: Revision of Mitrastema 223

Makino (1928), Matsuda ('Matuda') (1947), Van Royen (1963), and Hansen

(1973) have defended the opinion that there is only one.

Corner & Watanabe(1969) depicted what they considered to be 5 species, and

Corner (1978) could not identify the population from the Pinosok Plateau, Kinabalu,

Sabah, which is now destroyed, so suggesting it was an undescribed one.

Watanabe (1934d, 1936a) described geographic variation in Taiwan and Japan in

the numberof pairs of scales. The southern localities usually had 6 pairs but towards

the north there are only 4 or even 3. In Taiwan there were 8-12 pairs, while stems

with scales be correlation between the many are more angular. There appears to no

size of the flowers and the number of bracts. At some places flowers can be very

small, but the plants have many pairs of bracts.

kanehirae Intermediary forms between Mitrastema and M. yamamotoi occur in the

same area (Yamamoto, 1925; Watanabe, 1934d), while Van Royen (1963) reported

that the differences between M. kanehirae, M. kawa-sasakii, and M. yamamotoi were

foundto exist in the abundant New Guinea material, which undoubtedly represented

one species only.

whole cline in size and numberof of scales be Overall, a pairs seems to present,

and M. kanehiraeappears to be only an extreme form of M. yamamotoi.

the show in which is due the In Japan plants a seasonality flowering, to presence

of a dry season, in which they flower. Yamamoto distinguished a spring form

(M. yamamotoi) and an autumn form ( M. kanehirae). Palm (1934) observed that

in North Sumatra, where a dry season is absent, flowering occurs all around the

year (see Matsuda, 1947, for dataon mean temperature and rainfall in Japan, Taiwan,

and New Guinea Sumatra). In flowering seems to be seasonal, too (Van Royen,

1963).

One cause of the apparent variability is possibly due to edaphic and climatic fac-

tors. The depth of the roots of the hosts (Matsuda, 1947), how much litter is formed

in the of the forests, mean temperature, start dry season, moisture, day length, etc.

will doubt influence the host and the of its no development parasite. As some of

these factors are linked with latitude, the clines couldbe explained. Matsuda (1947)

suggested that the various forms are specific to their hosts. This seems an incorrect

assumption, see for instance the observations by Rao & Balakrishnan (1972) on the

where 4 different of Meghalaya population species 4 different families were being parasitised: Castanopsis tribuloides A. DC. (Fagaceae), Elaeocarpus lancifolius Roxb.

(Elaeocarpaceae), Engelhardtia spicata Blume (Juglandaceae), and Vernonia volka-

meriifolia DC. (Compositae).

that has been able Now WM to study more material than any previous student of the genus, we have come to the conclusion that the Southeast Asian specimens must be regarded as representatives of a single, variable species. This might seem to some

crude of but there a act lumping, are too many overlaps, gradual transitions, and too

in the characters. There be narrow gaps are, as might expected, local forms, or eco- types, and in due course some of these might evolve further, if they have not been eradicated before, but at present, without looking at the field label, it is often im-

with where collection and possible to say certainty a particular came from, so to what

'species' it would belong. 224 BLUMEA Vol. 38, No. 1, 1993

An example is the treatment ofthe Sumatrapopulation. Palm (1934) consideredit to

be conspecific with Mitrastemayamamotoi, Hayata (in litt. to Palm) identifiedit with

M. kawasasakii, but Nakai (1941a, b) based a new species on it, M. sumatranum.

The Yunnan form (Li, 1975) fits in well as a range extension from Cambodia and

Vietnam (Hansen, 1972, 1973).

Surprisingly, in 1924 a representative of the genus was discovered in the oak for-

ests of Chiapas, Mexico. It was described as a distinct species, M. matudae Y. Yama-

moto (1936, ‘matudai’), because of its much larger size, broadly triangular scales,

and triangular fruits, and its conical, more or less pyramidal stigma. Matsuda (1947)

gave a more extensive description, based on living material. At first sight this amphi-

distribution but there few similar of Pacific seems very strange, are quite a ones typi-

cal tropical genera. The explanation for such a curious disjunction remains unknown

(see Van Steenis, 1962, for a list more than 20 genera). Mitrastema matudae appears

distinct and be second for the quite must regarded as a species genus.

GENERAL LITERATURE ON MITRASTEMA

Airy Shaw, H.K. in J.C. Willis. 1973. A dictionary of the flowering plants and ferns: 750. Cambridge.

Akuzawa, E. 1985a. Problems concerning the period of occurrence of Kinabalu Mitrastemon. Para-

sitic PI. Mag. 13/4: 23-27.

Akuzawa, E. 1985b. Confirmation of the inflorescence of Mitrastemon in Mt. Kinabalu, North

Borneo. Parasitic PI. Mag. 14/2: 33-37.

Akuzawa, E. 1986. Blooming season of Kinabalu Mitrastemon. Parasitic PI. Mag. 15/1: 26-31.

Corner, E.J.H. 1978. In: D.M. Luping, C. Wen & E.R. Dingley (eds.), Kinabalu Summit of

Borneo. Sabah Soc. Monograph: 135-137. Kota Kinabalu, Sabah.

Corner, E.J.H., & K. Watanabe. 1969. Illustrated guide to tropical plants: 163. Tokyo.

Cronquist, A. 1981. An integrated system of classification of flowering plants: 703-704, t. 5, 35.

New York.

Gardner, G. 1844. In Hooker, Icon. PI. II, 3: t. 655.

1854. Plantae Gray, A. novae Thurberianae. Mem. Amer. Acad. Arts Sc., n.s. 2: 326, t. 7.

Hansen, B. 1972. Notes on some parasitic phanerogams from the Indo-Chinese Peninsula. Bot.

Tidsskr. 67: 146-151.

Hansen, B. 1973. Rafflesiaceae. Fl. Camb., Laos, Vietnam 14: 59-64, 9 fig.

Harms, H. 1935. Rafflesiaceae. In: H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. ed. 2,

16b: 243-281, t. 124-147. Leipzig, Berlin.

1913. liber Hayata, B. (23 Sept.) die systematische Stellung von Mitrastemon als einer neuenGat-

und besonderen tung Tribus der Rafflesiaceen. Bot. Jahrb. 51: 154-176, t. 1.

Hayata, B. (25 Dec.) 1913. On the systematic position of Mitrastemon. Icon. PI. Form. 3: 156,

199-213.

Li, H.-W. 1975. The Rafflesiaceae in China. Acta Phytotax. Sin. 13/2: 79-81, t. 1, f. 1-5.

Liu, T.-S., & M.-J. Lai. 1976. Rafflesiaceae. Fl. Taiwan 2: 582-584, t. 414, f. 1-4.

Mabberley, D.J. 1987. The Plant-book: 375. Cambridge, etc.

Makino, T. 1909. Mitrastemma yamamotoi. Bot. Mag. Tokyo 23: (325)—(327), fig.

Makino, T. 1911. Observations on the flora of Japan. Bot. Mag. Tokyo 25: 251-257, t. 7.

Makino, T. 1928. A contribution to the knowledge of the flora of Japan. J. Jap. Bot. 5: 18.

E. Matsuda, ('Matuda'). 1947.On the genus Mitrastemon.Bull. Torrey Bot. Club 74: 133-141.

Matsuura, H. 1932. (Tide in Japanese). Bot. Mag. Tokyo 46 (1932) 404-405. W. Meijer & J.F. Veldkamp: Revision of Mitrastema 225

Matsuura, H. 1935. A karyological investigation of Mitrastemon yamamotoi Mak. with special

in meiosis. J. Hokkaido Univ. Bot. 3: reference to the so-called 'diffuse' stage Fac. Sc. Imp. V,

189- 204, f. 5-8.

Nagai, K. 1934. Uber SagendaNagoya: Erklfirung von Nantozatsuwa.

Nakai, T. 1941a.Mitrastemon cochinchinensis Nakai. Icon. PI. As. Orient. 4: 306, t. 104.

Nakai, T. 1941b. Mitrastemon sumatranus Nakai. Icon. PL As. Orient. 4: 337-339, t. 113.

Okada, H. 1987. A Report of the Botanical Expedition to Papua New Guinea during 29th July and

2nd September, 1985. Sc. Rept., Col. Gen. Educ. Osaka Univ. 36/1: 15-17, fig.

Palm, B.T. 1934. A Mitrastemon, Rafflesiaceae, from Sumatra. Acta Horti Gotoburg. 9: 147-152,

fig-

Rao, A.S., & N.P. Balakrishnan. 1972. Mitrastemon in India. Ind. For. 98: 234-235, 2 fig.

Rose, J.N. 1909. Studies ofMexican and Central American plants. No. 6. Rafflesiaceae. The North

American species of Pilostyles. Contr. Nat. Herb. 12/6: 262-265, t. 20-23, 1 pi.

Rafflesiaceae. Guinea, 14: 243-245, 17. Royen, P. van. 1963. Sertulum Papuanum 8. Nova Bot. t.

Satake, Y„ et al. 1982. Wild Flowers of Japan 2: 110, t. 108.

Standley, P.C., & J. A. Steyermark. 1946. Fl. Guatemala. Rafflesiaceae. Fieldiana, Bot. 24: 101-104.

W.T. 1973. Botanical Latin, xiv 566 Newton Abbott. Stearn, + pp.

C.G.G.J, 1962. The in Blumea 11: 4-7. Steenis, van. land-bridge theory botany. 247-253, maps

Takhtajan, A.L, N.R. Meyer & V.N. Kosenko. 1985. Pollen morphology and classification in

Rafflesiaceae. Bot. Dzj. 70: 153-162, 4 tab.

1974. Mitrastemon found in North Borneo. J. Bot. 49: 214. Togashi, M. sp. Jap.

Watanabe, K. 1933a. Biologie von Mitrastemon yamamotoi Makino (Rafflesiaceae). I. Friichte und

Samen. Bot. Mag. Tokyo 67: 798-805, 3 fig.

Watanabe, K. 1933b. Ungeslechtliche Fortpflanzung von Mitrastemon yamamotoi. Proc. Imp. Acad.

Japan 9: 412-415, 2 fig.

Watanabe, K. 1934a. Geschlechdiche Fortpflanzung von Mitrastemon yamamotoi. Proc. Imp. Acad.

Japan 10: 421-423, f. 1.

Watanabe, K. 1934b. Biologie von Mitrastemon yamamotoi Makino (Rafflesiaceae). II. Vegetative

Fortpflanzung. Bot. Mag. Tokyo 48: 467-472.

Watanabe, K. 1934c. Uber die Flfissigkeitsabsonderung der Bliiten von Mitrastemon yamamotoi.

Proc. Imp. Acad. Japan 10: 507-508.

Watanabe, K. 1934d. Abweichende Formen von Mitrastemon yamamotoi. Proc. Imp. Acad. Japan

10: 600-603, t. 1-3.

Watanabe, K. 1935a. Beziehung zwischen Mitrastemon yamamotoi und M. kanehirai. Proc. Imp.

Acad. Japan 11: 280-282, t. 1.

Watanabe, K. 1935b. Mitrastemon kawa-sasakii aus Formosa. Proc. Imp. Acad. Japan 11: 283-

285, t. 1.

Watanabe, K. 1936a. Morphologisch-biologische Studien fiber die Gattung Mitrastemon I. J. Jap.

Bot. 12: 603-618.

Watanabe, K. 1936b. Morphologisch-biologische Studien Uber die Gattung Mitrastemon II. J. Jap.

Bot. 12: 698-711.

Watanabe, K. 1936c. Morphologisch-biologische Studien fiber die Gattung Mitrastemon III. J. Jap.

Bot. 12: 759-773.

Watanabe, K. 1936d. Morphologisch-biologische Studien Uber die Gattung Mitrastemon IV. J. Jap.

Bot. 12: 848-858.

Watanabe, K. 1937a. Morphologisch-biologische Studien Uber die Gattung Mitrastemon V. J. Jap.

Bot. 13: 14-24.

Watanabe,K. 1937b. Morphologisch-biologischeStudien Uber die Gattung Mitrastemon VI. J. Jap.

Bot. 13 : 75-86. 226 BLUMEA Vol. 38, No. 1, 1993

Watanabe,K. 1937c. Morphologisch-biologische Studien iiber die Gattung Mitrastemon VII. J. Jap.

Bot. 13: 154-162.

Yamamoto, Y. 1925. Species nova Rafflesiacearum ex Formosa. Bot. Mag. Tokyo 39: 142-145,

t. 1-15.

Yamamoto, Y. 1926. Rafflesiaceae. Suppl. Icon. Plant. Form. 2: 10-13. t. 5-8.

Yamamoto, Y. 1927. Notae ad plantas Japoniae et Formosae (VI). Mitrastemon Kawa-Sasakii

Hayata ex Sumatra. Bot. Mag. Tokyo 41: 736-737.

Yamamoto, Y. 1933. On Mitrastemon. Bot. & Zool. Tokyo 1: 1249-1258, t. 1-5, 1425-1428.

Yamamoto, Y. 1936. Species nova Mitrastemonacearum (Rafflesiacearum) ex Mexico. Bot. Mag.

Tokyo 50: 539-541, L 1-7.

MITRASTEMA

Mitrastema Makino, Bot. Mag. Tokyo 23 (1909) (326) l(‘Mitrastemma’). —Mitrastemon Makino,

Bot. Mag. Tokyo 25 (1911) 253, nom. inval. — Type species: Mitrastema yamamotoi Makino.

Parasites withoutchlorophyll, endoparasitic in roots, often inducing witches' brooms

often in around the its bark. there, gregarious, a fairy ring host, breaking through

Endophyte fungus-like. Basal part ofexophyte a corky, lenticellate, pusticular cupule, mainly composed of host tissue, rim irregularly 4-lobed. Stems unbranched,terete to

subquadrangular. Leaves sessile, scale-like, decussate, rarely whorled, in 3-7(-24) pairs, becoming larger upward, cream-coloured or milky yellowish-white at early flowering stage. Pedicels erect, terete or quadrangular. Flowers solitary, terminal, bi-

sexual, protandrous, hypogynous, actinomorphic. Perianth persistent, much reduced,

Stamens gamophyllous, collar-shaped, truncate to irregularly crenately 4-lobed. numerous, completely connate into a tube, widened at base, during early flowering

the of the but for small caducous stages surrounding apex pistil a aperture, during anthesis, removed by the growing pistil; apical part (connective) sterile, below this with several series of verticalrings with c. 10 anthers each; anthers minute, with extrorse slits. Pollen Pistil 1; longitudinal dicolpate. ovary superior, ellipsoid, 1-locular, apex half conical with a narrow transverse groove below the style; placentas radial,

8-15(-20), unequally deeply intruding; ovules numerous, anatropous, unitegmic, tenuinucellar; style 1, slightly constricted, stigma thick, conical, sub-bilobed. Fruit

with horizontal slit becoming a slightly woody, berry-like capsule, dehiscing a along the groove between the style and ovary. Seeds with a short funicle, sticky, testa hard, formed from the inner integument, yellowish to dark brown, reticulate because of collapsed cell walls when dry; endosperm a sweet jelly-like pulp, surrounding the embryo; embryo minute, undifferentiated, 4-celled.

Distribution - Species 2, one in Asia from India (Meghalaya) to S China (Yun- nan), Japan, Ryukyu I., Taiwan, and Malesia (Sumatra, Borneo, New Guinea), one in South America, from Mexico (Chiapas) to Guatemala and NW Colombia.

Habitat- Endophytic body living in the roots of older trees of species mainly of

Fagaceae (Castanopsis, Lithocarpus, Quercus, Trigonobalanus), infectedroots swol- len and growing near the surface with root witches' brooms. Endophytic body living like in the later included in older a mycelium, early stages near cambium, xylem. W. Meijer & J.F. Veldkamp: Revision of Mitrastema 227

Ecology - Seasonally flowering in subtropical areas, non-seasonally in tropical

Pollinated flies Palm, Van ones. by (Drosophila sp.: 1934; Royen, 1963), Chryso-

myia megacephala (F.), a bee (Apis sp.), wasps [Braconidae sp., Vespa magnified

nobilis Sonan (Matsuda, 1947) that crawl over the anthers], a firefly (Lampyridae

sp.) (Van Royen, 1963), and birds [e.g. Zosterops (palpebrosus) japonicus Tem-

& the excreted in the of the minck Schlegel, 1847] which feed on nectar axils upper-

most pairs of scales (Watanabe, 1934a). Seed dispersal most likely by these birds

and small mammals.

Chromosomes- x = 20 (Matsuura, 1935;Watanabe, 1936c).

KEY TO THE SPECIES

la. Flowers c. 6 cm long. Staminal tube 20-28 mm long; anthers in a 6-12 mm

broad ring. Ovary c. 16 by 15 mm; style c. 10 mm wide. Stigma 6-7 by 7-15

mm. - America 1. M. matudae

b. Flowers 1.5-2cm long. Staminal tube 14-20 mm long; anthers in a 2-5 mm

broad ring. Ovary c. 8 by 3 mm; style 3-6 mm wide. Stigma 2-5 by 4-6 mm.

- Asia 2. M. yamamotoi

1. Mitrastema matudae Y. Yamamoto

Mitrastema matudae Y. Yamamoto,Bot. Mag. Tokyo 50 (1936) 539 ('Mitrastemon matudai’). —

Lectotype: Matsuda 38595 (TI holo; MEXU), Mexico, Chiapas Prov., Soconusco Dist., Escu-

intla Municipality, Mt Ovando, 1700 m, parasitic onQuercus boqueronae Trelease, 5 April 1936.

Plants 4-12 3-3.5 1-1.5 lobed. Stems by cm. Cupule by 2-2.5 cm, margin weakly

c. 10 mm diameter.Leaves usually in 6 or 7 pairs, broadly to narrowly ovate, 13-30 by 10-25 mm, apex obtuse to apiculate. Flowers c. 6 cm long. Perianth 6-10 mm

Staminal 20-28 anthers in 6-12 high. tube mm long; a mm broad ring. Ovary ellip-

soid, c. 16 by 15 mm, style c. 2 by 10 mm, stigma 6-7 by 7-15 mm; placentas c. 15.

Fruit up to c. 10 by 15 mm. 2n = ? 63

Distribution - South America: Mexico (Chiapas Prov.), Guatemala (Alta Vera

Paz), Colombia(Dept. Antioquia).

Habitat- Montane rain or cloud forests, 1700-2500m altitude.

Ecology - Parasitic on Quercus boqueronae Trelease.

Collector's note - Bracts and total plant white.

2. Mitrastema yamamotoi Makino

Mitrastema Bot. 23 — Mitrastemon yamamotoi Makino, Mag. Tokyo (1909) (326), fig. yama-

motoi Makino, Bot. Mag. Tokyo 25 (1911) 255, t. 7. — Syntypes: Tashiro s.n., c. A° 1882,

Bando s.n., A° 1909, H. Yamamoto s.n., A° 1907, 1908, 1911 (TI), Japan, Kyushu.

Mitrastemon kawasasakii Hayata, Bot. Mag. Tokyo 26 (1912) 112, fig. — Mitrastemon yamamotoi

Makino kawasasakii J. Bot. 5 18. — Kawakami & var. Makino, Jap. (1928) Type: Sasaki s.n.

(TI holo), Taiwan, Monte Muto, 1250 m, Jan. 1911. 228 BLUMEA Vol. 38, No. 1, 1993

Mitrastemon kanehirae Y. Yamamoto, Bot. Mag. Tokyo 39 (1925) 142, f. 1-15 (''kanehirai’). —

Mitrastemon yamamotoiMakino var. kanehirae Makino, J. Jap. Bot. 5 (1928) 18 (‘kanehirai’).

— Type: Kanehira s.n. (TI holo), Taiwan, Rangeshi Nantou, on roots of Castanopsis jung-

huhnii (Miq.) Markgr. or Lithocarpus uraiana (Hayata) Hayata, Oct. 1924.

Mitrastemon cochinchinensis Nakai, Icon. PI. As. Or. 4 (1941) 306, t. 104. — Type: Anonymus

Institut des Recherches de l'lndochine s.n., Agronomique (TI holo), Cambodia,massif granite,

1700 m, Feb. 1930.

Mitrastema sumatranum Nakai [Icon. PL As. Or. 4 (1941) 306, nomen], Icon. PI. As. Or. 4 (1941)

338, t. 113. (‘Mitrastemon sumatranus’). — Type: Jochems s.n. (TI holo), Sumatra, Karo Mts,

1400 m, Aug. 1927.

Stems Plants 2.5-15 by 3 cm. Cupule 1-2 by 0.5-2 cm, margin irregularly lobed.

5-15 diameter. Leaves mm broadly to narrowly ovate, (4-)10-20(-28) by (5-)

10—15(—25) mm, apex bluntly rounded. Flowers 1.5-2 cm long. Perianth 5-10 by

6-17 mm. Staminal tube tubular, 14-20mm long; anthers in a 2-5 mm broadring.

Ovary ellipsoid, c. 8 by 3 mm; style ovoid-ellipsoid, 2-8 by 3-6 mm; stigma 2-5

= by 4-6 mm. Fruit up to c. 20 by 10 mm. 2n 40.

Distribution - India (Meghalaya), Thailand (Loei), Cambodia, Vietnam (Dalat),

China (Yunnan), Japan (Shikoku, Kyushu), Ryukyu I. (Amami I., Okinawa) (see

in Watanabe, Malesia: Sumatra E Borneo map 1937b), Taiwan; (Aceh, Coast), (Sabah: Kinabalu), Papua New Guinea (Western Highlands, Eastern Highlands,

Morobe, Milne Bay Prov.).

- 400-2000 Habitat In montane oak forests, m altitude.

Ecology - On Castanopsis tribuloides A.DC., Elaeocarpus lancifolius Roxb.,

Engelhardtia spicata Blume, and Vernonia volkameriifolia DC. in India (Rao & Bala- krishnan, elsewhere on C. acuminatissima (Blume) 1972); Castanopsis species, e.g. i

A. DC., C. cuspidata (Thunb.) Schottky and its var. sieboldii (Makino) Nakai, C.

taiwaniana Hayata, Eugenia sp., Ilex integra Thunb.,Limlia uraiana (Hayata) Masa-

& mune Tomiya, Lithocarpus elegans (Blume) Soepadmo, L. kawakamii Hayata,

Thunb., verticillata Quercus glauca Trigonobalanus Forman.Locally abundant, grow-

ing in large patches, up to 2 m diameter.

Collector's notes - Plant white, brownish white, pink, 5-7 cm long, stem cylin- drical to more or less quadrangular, c. 1 cm in diameter. Scales in 6-10 pairs, ob- conical, 1-2.5 by 0.8-2 cm, cream. Flowers with a fetid smell. Staminal column and ovary at first pink, later brown.

ACKNOWLEDGEMENTS

WM's research was partly financed with travel grants from the National Science Foundation, the

National GeographicSociety, and the University of Kentucky Research Foundation.

the both in the field and in the have WM in Over years many people, herbarium, helped finding

and conserved material. but be made living They are too many to mention, an exception must for

Dr. P.H. Raven who has moral and financial Dr. D.E. Breed- (MO) so long given strong support, love (CAS) who made a great effortto recollect and photographMeristema matudae in the field, and the staff members of the Kinabalu Park for their great help in the field.

Institutes visited their material loan for Many were or kindly sent on study, e. g. BM, BO, C,

US. All these that assisted CAS, K, L, M, MO, S, TI, TNS, and WM so much are herewith most gratefully acknowledged for their help, advice, and assistance. W. Meijer & J.F. Veldkamp: Revision of Mitraslema 229

SPECIMENS SEEN

(Only numbered or dated collections could be included)

— — 1909: — Boufford & Mitsuta 20128: Breed- Balakrishnan 34188: yam Bando A° yam (T) yam

love 25400: mat; 34366: mat.

Core 510: mat — Cruttwell 1645-A: yam.

Elliott 1052: yam.

— Harada — 12100: — Hatusima 11//1912: yam; 3/1937: yam Hartley yam 22806: yam Hayata

A° — & Ito — Honda 18/8/1954: 1915: yam Hayata 4/4/1914: yam 31/5/1939:yam; yam.

Jochems A° 1926: yam; 8/1927: yam (T).

Kanehira — Kanehira & Sasaki — Kawakami & Sasaki 10/1924: yam (T) 12/2/1918: yam 1/

— 1911: — Kimula Kinabalu Park Naturalist A° 1983: yam (T) 19/11/1921: yam yam.

Matsuda A° 1924 38595: — Muramatsu ('Matuda') (T): mat; mat (T) 25/8/1937: yam.

14579 & 30726 NGF (Womersley Millar): yam; (Streimann): yam.

Ohnuma — Ohwi & Akamoto 1482: 20/11/1913: yam yam.

1923: Palm A° yam. — A.S. Rao 37958: yam; 38222: yam; 38223: yam; 38224: yam; 38225: yam; 38285: yam van

11039: —RSNB 5746 & Royen yam; 15092: yam (Chew Comer): yam.

— 91455: — Streimann 8486: — Shingu 2/1913: yam Standley mat yam Suzuki 23/12/1931:

14502: yam; yam.

0 Tashiro A 1882: — 12/11/1912: — Tsai 58-8275: yam; 25/1/1924: yam Togashi yam yam.

Vink 16402: yam; 16402-A: yam.

— Wada al. 100: — Watanabe 27/2/1935: 2/1936: Womers- et yam 29/11/1934:yam; yam; yam

A° 1966: ley yam.

H. Yamamoto A° A 1908 A° 1911: — Y. Yamamoto al. 864: 1907: yam (T); (T): yam; yam (T) et

— Mori — Yamazaki 6981: yam Y. Yamamoto & 2-3/11/1932: yam yam; 26/12/1962:

yam; 9/1/1964: yam; 20/11/1978: yam.